Spatial Density Estimates of Eurasian Lynx (Lynx Lynx) in the French Jura and Vosges
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bioRxiv preprint doi: https://doi.org/10.1101/600015; this version posted August 4, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 1 Spatial density estimates of Eurasian lynx (Lynx lynx) in the French Jura and Vosges 2 Mountains 3 4 Olivier Gimenez1, Sylvain Gatti2, Christophe Duchamp3, Estelle Germain4, Alain Laurent2, 5 Fridolin Zimmermann5, Eric Marboutin2 6 7 1CEFE, CNRS, Univ Montpellier, Univ Paul Valéry Montpellier 3, EPHE, IRD, Montpellier, 8 France 9 2Office National de la Chasse et de la Faune Sauvage, ZI Mayencin, Gières, France. 10 3Office National de la Chasse et de la Faune Sauvage, Parc Micropolis, Gap, France. 11 4Centre de Recherche et d’Observation sur les Carnivores (CROC), 4 rue de la banie, 57590, 12 Lucy, France 13 5KORA, Thunstrasse 31, 3074, Muri, Switzerland 14 15 Abstract 16 17 Obtaining estimates of animal population density is a key step in providing sound 18 conservation and management strategies for wildlife. For many large carnivores however, 19 estimating density is difficult because these species are elusive and wide-ranging. Here, we 20 focus on providing the first density estimates of the Eurasian lynx (Lynx lynx) in the French 21 Jura and Vosges mountains. We sampled a total of 413 camera trapping sites (with 2 cameras 22 per site) between January 2011 and April 2016 in seven study areas across seven counties of 23 the French Jura and Vosges mountains. We obtained 592 lynx detections over 19,035 trap 24 days in the Jura mountain and 0 detection over 6,804 trap days in the Vosges mountain. Based 25 on coat patterns, we identified a total number of 92 unique individuals from photographs, bioRxiv preprint doi: https://doi.org/10.1101/600015; this version posted August 4, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 26 including 16 females, 13 males and 63 individuals of unknown sex. Using spatial capture- 27 recapture (SCR) models, we estimated abundance in the study areas between 5 (SE = 0.1) and 28 29 (0.2) lynx and density between 0.24 (SE = 0.02) and 0.91 (SE = 0.03) lynx per 100 km2. 29 We also provide a comparison with non-spatial density estimates and discuss the expected 30 discrepancies. Our study is yet another example of the advantage of combining SCR methods 31 and non-invasive sampling techniques to estimate density for elusive and wide-ranging 32 species, like large carnivores. While the estimated densities in the French Jura mountain are 33 comparable to other lynx populations in Europe, the fact that we detected no lynx in the 34 Vosges mountain is alarming. Connectivity should be encouraged between the French Jura 35 mountain, the Vosges mountain and the Palatinate Forest in Germany where a reintroduction 36 program is currently ongoing. Our density estimates will help in setting a baseline 37 conservation status for the lynx population in France. 38 39 Introduction 40 41 Obtaining estimates of animal population density is a key step in providing sound 42 conservation and management strategies for wildlife [1]. For many large carnivores however, 43 estimating density is difficult because these species are elusive and wide-ranging, resulting in 44 low detection rates [2]. To deal with these issues, non-invasive techniques, such as camera 45 trapping and DNA sampling, are increasingly used [3]. These non-invasive techniques 46 generate data that can be analyzed with capture-recapture methods to estimate densities [4]. 47 Standard capture-recapture models for closed populations [5] have long been used to 48 estimate animal abundance and density, including many large carnivores [6,7]. However, 49 when converting abundance into density, density estimates are highly sensitive to the size of 50 user-defined area assumed to reflect the effective sampling area [8]. In addition, individual bioRxiv preprint doi: https://doi.org/10.1101/600015; this version posted August 4, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 51 heterogeneity on the detection due to spatial variation in the distance of home ranges to the 52 sampling devices may lead to biased density estimates [5]. Spatial capture-recapture (SCR) 53 models deal with these issues by explicitly incorporating spatial locations of detections [9– 54 12], and they are increasingly used to estimate densities of large carnivores [13–18]. 55 Here, we focus on the threatened Eurasian lynx (Lynx lynx) in the French Jura and 56 Vosges mountains (see [19] for a map of its distribution in Europe; see also 57 https://www.lcie.org/Large-carnivores/Eurasian-lynx for recent updates). As in many regions 58 of western Europe [20], lynx were extirpated from France between the 17th and 20th centuries 59 due to habitat degradation, persecution by humans and decrease in prey availability [21]. 60 Shortly after their initial reintroduction in Switzerland in the 1970s [22], lynx naturally 61 increased their range and started recolonizing France by repopulating forests on the French 62 side of the Jura [21]. Reintroductions also occurred in the French Vosges mountain between 63 1983 and 1993 with the perspective of establishing a population there [23]. The species is 64 listed as endangered in the IUCN Red list and is of conservation concern in France due to 65 habitat fragmentation, poaching and collisions with cars and trains. Currently, the French 66 population of lynx is restricted to three mountain ranges, the Vosges, in northeastern France, 67 the Jura and the Alps, with little connectivity between them most likely due to human-made 68 linear infrastructures. While the Northern Alps are slowly being recolonized with lynx mostly 69 coming from the Jura [24], the Jura holds the bulk of the French lynx population. In contrast, 70 the lynx presence in the Vosges mountain remained stable following the reintroductions and 71 then, has been continuously decreasing since 2005 [25]. 72 Despite their conservation status, little information on abundance and density of lynx 73 in France exist. In this study, we used SCR and standard capture-recapture models to provide 74 the first estimate of lynx abundance and density using camera-trap surveys implemented in bioRxiv preprint doi: https://doi.org/10.1101/600015; this version posted August 4, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 75 the French Jura and Vosges mountains from 2011 to 2016. Based on these results, we discuss 76 research and management priorities for the effective conservation of lynx in France. 77 78 Methods 79 80 Ethics statement 81 We used non-invasive methods for data collection, which did not involve manipulation or 82 handling of any living organism. Therefore, approval from an animal ethics committee was 83 not required. Cameras were set on public or private forests with the permission of local 84 authorities or local owners, respectively. We advertised the study and the presence of camera 85 traps to the local stakeholders and the public visiting the areas. In agreement with French 86 legislation, we deleted photos permitting the identification of goods or people. 87 88 Study area and sampling design 89 The study area encompassed three counties of the French Jura mountain, namely Ain, Doubs 90 and Jura and four counties of the Vosges mountain, namely Vosges, Haut-Rhin, Bas-Rhin and 91 Moselle (Figure 1). Elevation ranged from 163 and 1,718 m above sea level in the Jura 92 mountain and from 104 to 1,422 m in the Vosges mountain. The human population density 93 was 88 per km2 in the Jura mountain and 170 per km2 in the Vosges mountain. Forests cover 94 50% on average of the Jura mountain [26] and 70% of the Vosges mountain [27]. Sampling 95 occurred over 6 years, between January 2011 and April 2016, mostly in winter and spring, 96 with surveys lasting between 2 and 4 months. We considered two study areas in 2011, 2014 97 and 2015, three study areas in 2013 and one study area in 2012 and 2016 through camera 98 trapping (Figure 1). 99 bioRxiv preprint doi: https://doi.org/10.1101/600015; this version posted August 4, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 100 [Figure 1 about here] 101 102 We divided each study area into a grid of 2.7 x 2.7 km cells applying a systematic 103 design where one out of two cells was sampled [28], hence ensuring that at least one camera 104 trap was set in each potential lynx home range (between 100km2 and 250km2, see [29]). To 105 maximize detectability, we set (non-baited) camera traps in forested habitats, based on 106 previous signs of lynx presence and on local knowledge, at optimal locations where landscape 107 and terrain features were likely to channel lynx movements on more predictable paths (on 108 forest roads, hiking trails and to a lesser extent on game paths) [30].